Reviewers: N/A
Editors: N/A

Initial release: 2019-08-23
Version information:

• Appearance: B
• Last minor update: 2022-01-22

Cite as: »Volstorf, Jenny. 2022. Cyprinus carpio (Dossier). In: fair-fish database, ed. fair-fish. World Wide Web electronic publication. Version B | 1.1. https://fair-fish-database.net.«

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• Unpredictable influence: no data found yet.
• Competition:
  • WILD: author concluded from previous research that Carp has high invasion potential because of attributes:
    a) successfully introduced on almost all continents, b) highly tolerant of temperature, salinity, pH, oxygen, c) genetically diverse (different strains), d) sexually mature at early age, e) fast succession of generations, f) growing rapidly, g) highly productive in terms of fecundity, h) opportunistic in diet, i) schooling, j) highly mobile, k) sought after by anglers (even if as bait). Climate matching suggests that Carp can survive almost everywhere in Australia. Comparison to other species showed that Carp were less affected by environmental changes, providing an advantage over competitors 1.
  • WILD: pairs of JUVENILES in cages (2.5 x 2.5 x 2 m depth) at 0.5 m depth in Utah lake containing macrophytes Potamogeton pectinatus, Ceratophyllum demersum, Scirpus validus, Chara aspera in 32 boxes per cage. After six weeks, lower macrophyte abundance and species diversity compared to control cages. Especially decrease in C. demersum and S. validus, no effect on P. pectinatus. Benthic macroinvertebrates decreased by 67%, especially amphipods, tabanids, hirudinids. Chironomids and oligochaetes increased probably because of newly exposed detritus.
    Second experiment to investigate effect of excluding Carp. Large exclosures (15 x 5 x 1.5 m) were placed on existing P. pectinatus beds at 1-1.25 m. After 30 days, 90% higher stem length of P. pectinatus and 58% more P. pectinatus compared to open cages that could be visited by Carp inhabiting the lake. Influence of lake side, though, with P. pectinatus decreasing despite excluded Carp probably due to missing protection from wind. No difference in invertebrates.
    Results of both experiments indicate indirect effect of Carp on macroinvertebrates by decreasing macrophyte abundance: Utah lake, Utah, (introduced) USA 2.
  • FARM/LAB, JUVENILES/ADULTS: Cyprinus carpio and Crayfish Cambarellus montezumae in experimental ponds (500 m², 1 m depth) to simulate situation in the wild where JUVENILES are stocked in irrigation ponds for crops at >2 IND/m². JUVENILES in experimental ponds at densities 0.3, 0.5, 1.7, or 3.8 IND/m² together with 70 Crayfish each. In experimental and irrigation ponds, decreasing Crayfish with increasing Carp density, with almost nonexistence from >0.4 IND/m² or catch per unit effort on. Diet in experimental and irrigation ponds consisted mostly of detritus in small (9-14 cm) and medium JUVENILES (18-23 cm), seeds and plant tissues in ADULTS (25-30 cm).
    In plastic tank (1.5 m diameter, 0.3 m height) divided in two segments by mesh material and filled with 350 g Potamogeton pectinatus, one small (10 cm) or large (35 cm) Carp with 23 Crayfish of different sizes that could swim to either side via plastic tubes (5 cm diameter). After three days of three times two hours exploration by Crayfish each, Crayfish preferred sections with plants and avoided section with Carp (even if containing plants). Results indicate that Carp displace Crayfish not by predation but by separating them from macrophytes and probably also by increasing turbidity when foraging through sediment which in turn decreases macrophyte density that Crayfish depend on 3.
  • LAB: authors concluded from previous research that Carp – through foraging behaviour – resuspends bottom sediments which leads to a) increased turbidity, nutrients, phytoplankton, b) decreased invertebrate density and diversity, c) increased uprooting of macrophytes. Effect "a" results in i) shading and thus decreased macrophyte growth, ii) blocking of habitat and thus decreased invertebrate density, iii) altering community structure and thus decreased zooplankton. Ecosystem changes from oligotrophic (clear) to eutrophic (turbid) and decreases abundance of amphibians, waterfowl, fishes 4.
• Disease transmission: no data found yet.
• Interbreeding:
  • WILD: hybridisation between C. carpio and also invasive Carassius auratus. Probably increases invasive potential due to enhanced genetic diversity, for example by introducing resistance to Koi herpes virus via C. auratus: Murray-Darling-Basin, Australia (introduced) 5.

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• For feeding ➝ 6.
• For daily rhythm ➝ 7.
• For swimming ➝ 8 9 10 11 7.
• For courtship, spawning ➝ 12.
• For aggregating ➝ 8.
• For memory ➝ 7.

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• For feeding ➝ 13 14.
• For daily rhythm ➝ 15.
• For courtship and spawning ➝ 16.
• For learning ➝ 17 15 18.
• For coping styles ➝ 19 20 21 22 18.
• For observable stress reactions ➝ 23 24 25 26 27.

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• Observations Danube river: 29.

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• Observations Africa: Amerti reservoir, Ethiopia 30, Hardap reservoir, Namibia 9, lake Gariep, South Africa 31.
• Observations Australia: Barwon river, Victoria, Australia 32, Broken river (Murray-Darling-Basin), Australia 10, Campaspe irrigation district, Victoria, Australia 32, Gippsland lakes, Victoria, Australia 32, lake Dartmouth, Victoria, Australia 32, lake Eildon, Victoria, Australia 32, lake Modewarre, Victoria, Australia 32, Murray-Darling-Basin, Australia 5, Murray river, Victoria, Australia 33 32, Wimmera river, Victoria, Australia 32.
• Observations Europe: Almus dam lake, Turkey 34, Cernek lake, Turkey 35, Gölhisar lake, Turkey 36, estuary of Guadalquivir river, Spain 37, lake Banyoles, Spain 38, Liman lake, Turkey 39.
• Observations New Zealand: Lower Waikato region, New Zealand 40.
• Observations North America: Acambay valley, Mexico 3, Clear lake, Iowa, USA 11, lake Erie, Canada/USA 8, lake Susan, Minnesota, USA 7, Marsh and Pelican lake, Nebraska, USA 41, overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA 12, Utah lake, Utah, USA 2.

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• Observations African catfish WILD: Clarias gariepinus: Hardap reservoir, Namibia (introduced) 9.
• Observations Black Bullhead WILD: Ameiurus melas: Marsh lake and Pelican lake, Nebraska, USA (introduced) 41.
• Observations Bluegill WILD: Lepomis macrochirus: Pelican lake, Nebraska, USA (introduced) 41.
• Observations Chub WILD: Leuciscus cephalus: lake Banyoles, Spain (introduced) 38.
• Observations Common rudd WILD: Scardinus erythrophthalmus: Cernek lake, Turkey (introduced) 35.
• Observations Crucian carp WILD: Carassius carassius: Liman lake, Turkey (introduced) 39, Cernek lake, Turkey (introduced) 35.
• Observations Dnieper chub WILD: Leuciscus borysthenicus: Gölhisar lake, Turkey (introduced) 36.
• Observations Flathead grey mullet WILD: Mugil cephalus: Liman lake, Turkey (introduced) 39, Cernek lake, Turkey (introduced) 35.
• Observations Freshwater blenny WILD: Salaria (= Blennius) fluviatilis: lake Banyoles, Spain (introduced) 38.
• Observations Golden shiner WILD: Notemigonus crysoleucas: Marsh lake, Nebraska, USA (introduced) 41.
• Observations Largemouth bass WILD: Micropterus salmoides: lake Banyoles, Spain 38, Pelican lake, Nebraska, USA (introduced) 41.
• Observations Moggel WILD: Labeo umbratus: Hardap reservoir, Namibia (introduced) 9.
• Observations Mosquitofish WILD: Gambusia affinis: Gölhisar lake, Turkey (introduced) 36.
• Observations Mozambique tilapia WILD: Oreochromis mossambicus: Hardap reservoir, Namibia (introduced) 9.
• Observations Mudfish-Moggel hybrid WILD: Labeo capensis x Labeo umbratus hybrid: Hardap reservoir, Namibia (introduced) 9.
• Observations Nile tilapia WILD: Oreochromis niloticus: Amerti reservoir, Ethiopia (introduced) 30.
• Observations Northern pike WILD: Esox lucius: Pelican lake, Nebraska, USA (introduced) 41.
• Observations Orange river mudfish WILD: Labeo capensis: Hardap reservoir, Namibia (introduced) 9.
• Observations Pikeperch WILD: Sander lucioperca: Liman lake, Turkey (introduced) 39, Cernek lake, Turkey (introduced) 35.
• Observations Pumpkinseed sunfish WILD: Lepomis gibbosus: lake Banyoles, Spain (introduced) 38.
• Observations Redbelly tilapia WILD: Tilapia zillii: Amerti reservoir, Ethiopia (introduced) 30.
• Observations Roach WILD: Rutilus rutilus: lake Banyoles, Spain (introduced) 38.
• Observations Squalius cii WILD: Leuciscus cephalus: Cernek lake, Turkey (introduced) 35.
• Observations Straightfin barb WILD: Barbus paludisnosus: Hardap reservoir, Namibia (introduced) 9.
• Observations Toothcarp WILD: Aphanius sp.: Cernek lake, Turkey (introduced) 35.
• Observations Wels catfish WILD: Silurus glanis: Gölhisar lake, Turkey (introduced) 36.
• Observations Yellowfish WILD: Barbus cf. kimberleyensis (B. aenus x B. kimberleyensis hybrid): Hardap reservoir, Namibia (introduced) 9.
• Observations Yellow perch WILD: Perca flavescens: Marsh lake and Pelican lake, Nebraska, USA (introduced) 41.
• Observations Zander WILD: Stizostedion lucioperca: Gölhisar lake, Turkey (introduced) 36.

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• Plants:
  • WILD: FRY were found in the vicinity of bog-rush (Juncus), algae, pondweed (Potamogeton), JUVENILES near hornwort (Ceratophyllum): overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12.
  • WILD: JUVENILES were caught between reed and submerged macrophytes: river Murray, Australia (introduced) 6.
  • WILD: JUVENILES-ADULTS mainly stayed in areas with dead trees: Hardap reservoir, Namibia (introduced) 9.
  • WILD: JUVENILES-ADULTS preferred beds of emergent vegetation, mainly giant bulrushes (Scirpus validus) and cattails (Typha spp.), to other substrate types: Clear lake, Iowa, USA (introduced) 11.
  • For substrate and spawning ➝ D1.
• Rocks and stones:
  • WILD: JUVENILES-ADULTS mainly stayed in areas with gravel: Hardap reservoir, Namibia (introduced) 9.
  • WILD: JUVENILES-ADULTS avoided rocky areas: Clear lake, Iowa, USA (introduced) 11.
• Sand and mud:
  • WILD: JUVENILES-ADULTS were found over sandy bottom, mud: lakes in vicinity of Thompson's lake, Illinois, USA (introduced) 12.
  • WILD: JUVENILES-ADULTS mainly stayed in areas with sand and mud: Hardap reservoir, Namibia (introduced) 9.
  • WILD: individuals were found over sand and mud: Utah lake, Utah, USA (introduced) 2.
  • WILD: JUVENILES-ADULTS were observed over silt and sand, especially in shallow water and spring: Clear lake, Iowa, USA (introduced) 11.
• Other substrate: no data found yet.

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• Plants:
  • WILD: FRY were found in vegetation where they were spawned (for spawning substrate ➝ D1), not in open water. Were observed feeding among moss: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12.
• Rocks and stones: no data found yet.
• Sand and mud: no data found yet.
• Other cover: no data found yet.

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• Observations: 3.1±0.0 se 42.

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• Omnivorous D2 The fishery that provides fish meal and fish oil has two major impacts:
  1. It contributes considerably to overfishing, as it accounts for 1/4 43 or even 1/3 44 of the world catch volume.
  2. It challenges animal welfare, because in the face of 450-1,000 MILLIARD wild fishes caught worldwide each year to be processed into fish meal or fish oil 45, the individual fish gets overlooked and, thus, suffering increases at rearing, live marketing, and slaughtering levels 46.

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• Food items: omnivorous:
  • Observations WILD, FRY: copepods (Cyclops), cladocerans (Alona), ostracods, amphipods (Hyalella knickerbockeri): overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12.
  • Observations WILD, JUVENILES-ADULTS: detritus, plant material (debris, diatoms, seeds), amphipods (Echinogammarus spp.), phantom midge larvae (Chaoborus flavicans): lake Banyoles, Spain (introduced) 38.
• Food items and habitat:
  • LAB: JUVENILES in rectangular glass tanks (2.5 x 0.4 x 0.9 m) at density 3 IND/tank with either a) plankton (P tanks), b) plankton and benthic macroinvertebrates with 100 mm pond sediment (PB tanks), or c) plankton, benthic macroinvertebrates, and artificial feed (at 10:00 h) with 100 mm pond sediment (PBF tanks). During four weeks, increase in phyto- and zooplankton in PB and PBF tanks, more so in PBF; in P tanks, stagnation in phytoplankton, decrease in zooplankton. Results probably due to mainly feeding on macroinvertebrates in PB (76%) and PBF tanks (65%), mainly on zooplankton (71%) in P tanks. Highest specific growth rate in PBF (ca 2.5%/d) compared to PB (1.5%/d) compared to P tanks (1.0%/d). Available food influenced feeding position: in P tanks, mainly feeding in water column, in PBF tanks, mainly on bottom. Available food also influenced social behaviour: in P tanks, more dispersed grazing. No aggressive behaviour 13.
• Food items and life stages: mainly aquatic invertebrates, increasing in size with increasing age:
  • WILD: FRY mainly cladocerans (Daphnia sp.), JUVENILES mainly cladocerans, but also sand, Phragmites sp. seeds, ostracoda: river Murray, Australia (introduced) 6.
  • WILD: smaller individuals preyed on meiobenthos (cladocerans, ostracods), larger on profundal macrobenthos (phantom midge larvae, large chironomids): lake Banyoles, Spain (introduced) 38.
• Food preference:
  • LAB: JUVENILES in 45 L tanks at density 3 IND/tank acclimatised to 16 °C. After 52 days, gradually changed to 20 °C, 24 °C or 28 °C by increasing temperature by 0.5 °C/d. During 50 days, at each temperature, preferred pelletised fish meal (2 mm) over maggots 14.
• Food partitioning:
  • WILD: great overlap in diet of introduced C. carpio and Rutilus rutilus in terms of detritus, plant material, amphipods, but in R. rutilus, cladoceran Daphnia longispina, nematoceran pupae Chaoborus flavicans, and exuviae played larger role. In C. carpio, plant seeds and hard debris, mollusks (Physella acuta, Pisidium spp.), ostracods (Candona sp., C. ophtalmica) were more important: lake Banyoles, Spain (introduced) 38.
• Prey density:
  • LAB: JUVENILES in 45 L tanks acclimatised to 16 °C and gradually changed to 20 °C, 24 °C or 28 °C by increasing temperature by 0.5 °C/d were presented with 20, 30, 40, 50, 70, 100, or 150 food items/tank equalling 148, 222, 296, 370, 519, 740, 1,111 items/m². Higher feeding rate the higher the temperature and the higher the prey density 14.
• Prey size selectivity: no data found yet.
• Particle size: no data found yet.

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• LAB: FRY were fed a diet with 1, 2, or 3% fructo-oligosaccharide. After seven weeks, higher survival rate (98.3% versus 61.6%) and higher leucocyte count (39.3 x 10³ per µL versus 32.6 x 10³ per µL) under 3% supplementation compared to control, 1% and 2% supplementation in between (78.4-83.3%, 36.3-37.3 x 10³ per µL). No difference in erythrocyte count, haematocrit. Higher level of heterotrophic autochthonous gut bacteria (5.6-5.7 colony forming units/g versus ca 4.8 under 1% versus 4 in control) and autochthonous lactic acid bacteria (2.4 colony forming units/d under 3% versus 1.5 under 2% versus 0 under 1% and control) under 2 and 3% supplementation than 1% and control. FRY subjected to 15 g/L salinity stress test had higher survival when fed fructo-oligosaccharides compared to control (53.4-59.6% versus ca 35%) 49.

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• WILD: large quantity of sand in stomachs of JUVENILES indicates bottom-feeding: river Murray, Australia (introduced) 6.
• For feeding style and available food items ➝ D2.

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• LAB: JUVENILES in 150 L glass tanks with pendulum rod at 10 cm below water surface connected to self-feeder. During 20 days, higher glucose at 22:00 h (peak of self-feeding activity) than 10:00 h (nadir), but lower levels compared to pair-fed control (22:00 h: ca 2.5 mmol/L versus 3.5 mmol/L, 10:00 h: ca 1.5 mmol/L versus 2 mmol/L). No difference in cortisol levels in self-feeding JUVENILES between 22:00 h and 10:00 h compared to increase at 10:00 h (feeding time) in control JUVENILES (22:00 h: 1.5 nmol/L versus 2.2 nmol/L, 10:00 h: 3.3 nmol/L versus 130 nmol/L) probably due to anticipation of food 15.

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• LAB: JUVENILES in 150 L glass tanks with pendulum rod at 10 cm below water surface connected to self-feeder. During 20 days, JUVENILES activated the self-feeder on average one time per individual per day (with peak at 22:00 h), amounting to food at 4% body weight. After 20 days, higher weight (46.5 g versus 39.9 g), higher body weight increase (156 g/20 IND versus 31 g/20 IND), and higher specific growth rate (0.8% versus 0.2%) compared to controls fed similar ration by hand once a day at 10:00 h. Result is possibly due to synchrony in self-feeding individuals with biorhythm and highest gut activity 15.

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• Feeding and temperature:
  • LAB: JUVENILES in 45 L tanks at density 3 IND/tank acclimatised to 16 °C. After 52 days gradually changed to 20 °C, 24 °C or 28 °C by increasing temperature by 0.5 °C/d. After 50 days, higher feeding rate the higher the temperature with no difference between 24 °C and 28 °C (ca 1.2-1.3 food items/s under 24 °C and 28 °C versus ca 1-1.1 items/s under 20 °C versus ca 0.8-1.1 items/s) 14.

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• Daily rhythm:
  • WILD: ADULTS visited mesh bag of cracked corn placed in lake along shoreline mostly from dusk till dawn, peaking at 22:00 h: lake Susan, Minnesota, USA (introduced) 7.
  • LAB: JUVENILES in rectangular glass tanks (2.5 x 0.4 x 0.9 m) at density 3 IND/tank grazed mainly during the day (43% during day versus 26% at night), swam mainly at night (73% at night versus 56% during day). Peak of feeding activity at 08:00 h and 17:00 h. Schooling (i.e., at least two JUVENILES together) at night, dispersed grazing during the day 13.
  • LAB: JUVENILES activated self-feeder at higher frequency during night than day with peak at 22:00 h and nadir at 10:00 h 15.
• Nocturnal activity: ➝ Daily rhythm.
• Phototaxis: no data found yet.
• For daily rhythm and home range ➝ D7.

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• LAB: JUVENILES (Scaled carp, 50.5 g) in 147 L glass tanks (100.5 x 40 x 36.5 cm) or a recirculating aquaculture system at density 10 IND/tank, either 80 or 200 lux and either Mozart's "Eine kleine Nachtmusik" or "Romanza - Jeux interdit" (anonymous) during 4 h per day Monday-Friday or no music. Ambient noise level at 123 dB re 1 µPa, music up to 130 dB re 1 µPa. After 106 days, lower haematocrit under 200 than 80 lux (24.9-25.1 versus 26.6-27.9). No difference in glucose (64.9-74.5 mg/dL) and various brain neurotransmitters (DOPAC, HVA, 5-HIAA) 50.

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• LAB: JUVENILES (Scaled carp, 50.5 g) in 147 L glass tanks (100.5 x 40 x 36.5 cm) or a recirculating aquaculture system at density 10 IND/tank, either 80 or 200 lux and either Mozart's "Eine kleine Nachtmusik" or "Romanza - Jeux interdit" (anonymous) during 4 h per day Monday-Friday or no music. Ambient noise level at 123 dB re 1 µPa, music up to 130 dB re 1 µPa. After 106 days, higher weight (156.7-177 g versus 154.1-161.8 g), higher specific growth rate (1.07-1.18% versus 1.05-1.1%), and lower FOOD CONVERSION RATIO (1.88-2.01 versus 1.98-2.03) under 200 lux than under 80 lux but only in music groups not in control, with the best values under Romanza (177 g, 1.18%, 1.88) 50.

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• Standard temperature range: 0.1-29.3 °C, in extreme cases up to 33 °C:
  • Observations WILD: FRY in water holes of almost dried up fields and marshes in 31-33 °C: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, 2-14 °C: warm-water discharge canal, lake Erie, Canada/USA (introduced) 8, 8-27 °C: Gölhisar lake, Turkey (introduced) 36, 7-26 °C: lake Banyoles, Spain (introduced) 38, 12-28 °C: Hardap reservoir, Namibia (introduced) 9, 9-18 °C April-July, 10-25 °C August-March: various sampling sites, Victoria, Australia (introduced) 32, 8-25 °C: Broken river (Murray-Darling-Basin), Australia (introduced) 10, 21.3-29.3 °C: Utah lake, Utah, USA (introduced) 2, Koi carp: 11-23 °C: Lower Waikato region, New Zealand (introduced) 40, 3 °C water surface temperature in winter, 10 °C in spring and autumn, 25.5 °C in summer, mean 17.4 °C: Almus dam lake, Turkey (introduced) 34, 0.1-25.1 °C: Clear lake, Iowa, USA (introduced) 11, 15-27 °C: lake Susan, Minnesota, USA (introduced) 7, 8.7-11.3 °C in winter, 16.6-26 °C in summer: lake Gariep, South Africa (introduced) 31, 18.9-23.1 °C: Amerti reservoir, Ethiopia (introduced) 30.
• Temperature preference:
  • WILD: individuals (420-630 mm, 1,158-3,671 g) were resident in warm-water discharge canal (length 550 m, width 15.3 m, depth 9.1 m, 2-14 °C) from thermal-generating power station during winter. Residency in canal decreased throughout spring with increasing temperatures (residency 100% in December versus ca 30% in April) in adjacent lake Erie, Canada/USA (introduced) 8.
• Migration temperature: no data found yet.
• For temperature and...
  ...depth ➝ D8,
  ...spawning ➝ D1.

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• Lower and upper lethal limits: no data found yet.
• Temperature (change) and stress:
  • WILD: individuals (41.1-78 cm) tagged internally (compared to externally) did not survive until the end of the observation period, probably because more vulnerable to handling and surgery due to high temperatures (24-25 °C) at time of tagging: Hardap reservoir, Namibia (introduced) 9.
  • LAB: eggs from hormonally stimulated 7-8 kg females were fertilised with milt from hormonally stimulated 5-8 kg males at 22-23 °C and transferred to 20 °C, 24 °C, 28 °C, or 32 °C immediately. Survival to stage of swim bladder inflation was 76.6%, 80%, 64.2%, 0% 51.

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• Temperature must exceed: no data found yet.
• Temperature must not go beyond: no data found yet.
• Optimal temperature for growth:
  • LAB: eggs from hormonally stimulated 7-8 kg females were fertilised with milt from hormonally stimulated 5-8 kg males at 22-23 °C and transferred to 20 °C, 24 °C, 28 °C, or 32 °C by changing temperature at 2-3 °C/h. With beginning of exogenous feeding, FRY were transferred to 20 dm³ aquaria at density 400 IND/tank and water changed to 23 °C at rate of 2-3 °C/h. Faster development the higher the temperature, e.g. later hatching (3.2-4.7 days versus 1.5 days), later inflation of swim bladder (3.8-6.5 days versus 2.4-2.5 days), and later beginning of exogenous feeding under 20 °C compared to 32 °C (5.6-7.1 days versus 2.8-3.2 days). Shorter length the higher the temperature, e.g. at hatching (5.7 mm at 20 °C versus 5.1 mm at 32 °C). After 14 days at 23 °C, FRY with highest total length (19.1-21.2 mm versus 18.1-18.7 mm) and highest weight (92.9-178.7 mg versus 83.4-108.1 mg) in former 24 °C- or 28 °C-groups compared to 20 °C- or 32 °C-groups. No difference in survival since fertilisation except for more variation in one experiment which decreased after hatching (84.6-89.1%). Results indicate optimal temperature for survival is 20-28 °C, for growth is 24-28 °C, and for development is 28-32 °C 51.
  • LAB: JUVENILES in 45 L tanks at density 3 IND/tank acclimatised to 16 °C. After 52 days gradually changed to 20 °C, 24 °C or 28 °C by increasing temperature by 0.5 °C/d. After 50 days, higher specific growth rate under higher temperature compared to 16 °C (0.5-0.6%/d versus 0.3-0.4%/d) with highest growth under 20 °C 14.
• For temperature and feeding ➝ D9.

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• Salinity tolerance:
  • Natural and introduced distribution in fresh water or brackish water the most D10 D11.
• Standard salinity range:
  • Observations WILD: average 1.5 g/L: Utah lake, Utah, USA (introduced) 2, 2-4 ppt: Liman lake, Turkey (introduced) 39, average 0.8-1.3 ppt: Cernek lake, Turkey (introduced) 35.

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• Standard turbidity range:
  • Observations oligo-mesotrophic WILD: lake Gariep, South Africa (introduced) 31.
  • Observations mesotrophic WILD: lake Banyoles, Spain (introduced) 38.
  • Observations eutrophic WILD: Clear lake, Iowa, USA (introduced) 11.
• Secchi depth (water transparency): 5-49 cm:
  • Observations WILD: 5-14 cm: Hardap reservoir, Namibia (introduced) 9, 12 cm Secchi depth: Utah lake, Utah, USA (introduced) 2, <0.5 m: lake Susan, Minnesota, USA (introduced) 7.
• For turbidity and...
  ...escapees ➝ D12,
  ...feeding style ➝ D13.

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• WILD: two individuals (33-49 cm) were brought to release site ca 4,500 m from catch site. Both returned to catch site. Range of daily distance moved of all five radio-tagged individuals was 0.003-10.6 km/d, amounting to 0.03-4.0 km². Total area utilised during four months observation period ranged at 1.5-7.6 km², mean 4.3 km²: Hardap reservoir, Namibia (introduced) 9.
• WILD: during late summer (February) to mid winter (July), four of five individuals displayed site fidelity around release site and established home range of 10-50 m core area with occasional visits of up to 860 m, the last individual moved 2,800 m away and established home range there. Ten individuals were brought to release site 2,250-2,710 m upstream or 1,870-2,720 m downstream from capture site in plastic box. Of the four ones translocated upstream, two returned to capture site and established home range, one established home range at release site, one ca 800 m upstream of release site. Of the five ones translocated downstream, three returned to capture site and established home range, one established temporary home range at release site, one ca 1,000 m downstream of release site. Core area of home range of translocated individuals ranged around 10-70 m with occasional visits up to 600 m. Results indicate site fidelity but also shifts of home ranges: Broken river (Murray-Darling-Basin), Australia (introduced) 10.
• WILD, ADULTS: daily core area of distribution during day of average 67,891 m² (0.07 km²), during night of 70,614 m² (0.07 km²): lake Susan, Minnesota, USA (introduced) 7.

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• Depth range in the wild:
  • Observations WILD, FRY-ADULTS: FRY: 15-120 cm: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, individuals (345-614 mm fork length) 0-20 m: lake Banyoles, Spain (introduced) 38, individuals (33-49 cm) mainly stayed in open areas with 5-25 m and shallow inlets of 1-5 m: Hardap reservoir, Namibia (introduced) 9, JUVENILES mean 0.7-1.3 m during spring and autumn, ADULTS mean 1.5-3.2 m: Clear lake, Iowa, USA (introduced) 11.
  • For depth and spawning ➝ D1.
• Depth in cages or tanks: no data found yet.
• Depth preference: <1 m:
  • Observations WILD, JUVENILES-ADULTS: individuals (420-630 mm, 1,158-3,671 g): <1 m: warm-water discharge canal, lake Erie, Canada/USA (introduced) 8.
• Depth and daily rhythm: no data found yet.
• Depth and low temperatures:
  • WILD: individuals (345-614 mm fork length) were caught more in the littoral in spring: lake Banyoles, Spain (introduced) 38.
  • WILD: ADULTS moved deeper in autumn (mean 2.2 m, mean 9.3 °C) and winter (mean 2.3 m, mean 0.1 °C) than in spring (mean 1.7 m, mean 12.4 °C) and summer (mean 1.6, mean 24.3 °C): Clear lake, Iowa, USA (introduced) 11.
• Depth and high temperatures:
  • WILD: individuals (345-614 mm fork length) were caught deeper in winter than in spring: lake Banyoles, Spain (introduced) 38.
• Position in habitat and age: no data found yet.
• Depth and light intensity: no data found yet.
• Depth and noise: no data found yet.
• Depth and threat: no data found yet.

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• JUVENILES and ADULTS migrate within fresh water, ADULTS migrate to spawning grounds:
  • For migration and...
    ...water temperature ➝ D14,
    ...home range ➝ D7,
    ...spawning ➝ D1.

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• Observations time from fertilisation until hatching WILD: 2-3 days (23 °C): Danube river 29.
• Observations time from fertilisation until hatching LAB: 60-90 h depending on temperature (faster the warmer) 16, 48 h 52, 3.2-4.7 days at 20 °C, 1.5 days at 32 °C 51.
• Observations size WILD: 0.8-2.5 mm diameter: estuary of Guadalquivir river, Spain (introduced) 37, 1.5-1.8 mm diameter: Danube river 29, 893-1,772 µm: various sampling sites, Victoria, Australia (introduced) 32, Koi carp: 0.3-1.9 mm mean yolked oocyte diameter: Lower Waikato region, New Zealand (introduced) 40.
• Observations size FARM: 1.1-1.3 mm 53.
• Observations size LAB: 1.4-2 mm diameter 16, 1.9 mm diameter 51.
• Observations weight: no data found yet.

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• Observations yolk sac absorption LAB: day 4-7, 7-9.5 mm 16.
• Observations age and TOTAL LENGTH LAB: 3.9-5.6 mm at hatching 16, 4.5 mm at hatching, 12 days: 12 mm 52, 5.7 mm at hatching at 20 °C, 5.1 mm at hatching at 32 °C 51.

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• Observations age at beginning of exogenous feeding WILD: 2 days: Danube river 29.
• Observations age at beginning of exogenous feeding LAB: 24-36 h after hatching 16, 5.6-7.1 days at 20 °C, 2.8-3.2 days at 32 °C 51, 5-7 days 55.
• Observations age and TOTAL LENGTH WILD: 12-15 days, 6.4-12.7 mm: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, 5-29 days, 7.1-16.7 mm standard length: river Murray, Australia (introduced) 6.
• Observations TOTAL LENGTH and weight LAB: 7.8-16 mm 16, 45 days, 26.8 mm standard length, 0.6 g 56, 3.2 g 49.

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• Observations age, TOTAL LENGTH, and weight WILD: 31-61 days, 16.6-57.3 mm standard length: river Murray, Australia (introduced) 6, 150-193 mm, 51 g 2, Koi carp: 1 year, range 100-119 mm fork length: Lower Waikato region, New Zealand (introduced) 40, 203-330 mm, 150-490 g: Clear lake, Iowa, USA (introduced) 11.
• Observations age, TOTAL LENGTH, and weight FARM: 2 years and 192-200 g 57, 5.3 cm 3.
• Observations age, TOTAL LENGTH, and weight LAB: 16-18 mm, first scales at 17-18 mm, complete scale cover at 24-30 mm 16, 1 year, 9-12 cm, 12-15 g 58, 30 days and 25 mm 52, 270 days and 101.7 g, 300 days and 99.5 g 59, 160 days and 85.1 g 60, 33.1 g 23, 90.3-98.7 g 13, 64.7 mm, 1 year and 25.3 g, 1 year and 64.7 g 19, <1 year, 20-57 g 20, 3.7 g 61, 1 year, 90-105 mm fork lenght, 21.5 g 14, 38.5 g 15, 11.3-12.8 cm standard length, 40-64 g 62, 8.4 cm, 10.1 g 18.
• Sexual maturity for 50% of juveniles: males: 2-3+ years, 312 mm or less, females: 2-3+ years, 312+ mm:
  • Observations age and TOTAL LENGTH of males WILD: both sexes matured at 2+ years: estuary of Guadalquivir river, Spain (introduced) 37, Koi carp: at 2 years: Lower Waikato region, New Zealand (introduced) 40, 100% maturity of both sexes by 360 mm, 50% by 3.4 years and 312 mm, smallest mature male: 171 mm: Marsh lake, Nebraska, USA (introduced) 41, 50% maturity by ca 2 years, 295.9 mm fork length, smallest mature male: 258 mm fork length: lake Gariep, South Africa (introduced) 31, 50% sexual maturity at 27.2 cm fork length, smallest 15.5 cm fork length and 74.3 g: Amerti reservoir, Ethiopia (introduced) 30.
  • Observations age and TOTAL LENGTH of females WILD: both sexes matured at 2+ years: estuary of Guadalquivir river, Spain (introduced) 37, Koi carp: at 3 years: Lower Waikato region, New Zealand (introduced) 40, 100% maturity of both sexes by 360 mm, 50% by 3.4 years and 312 mm, smallest mature female: 348 mm: Marsh lake, Nebraska, USA (introduced) 41, 50% maturity by ca 2 years, 334.8 mm fork length, smallest mature female: 276 mm fork length: lake Gariep, South Africa (introduced) 31, 50% sexual maturity at 28.3 cm fork length in females, smallest 17 cm fork length and 86.9 g: Amerti reservoir, Ethiopia (introduced) 30.

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• Observations age, TOTAL LENGTH, and weight WILD: 6 years and 438.2 mm fork length: estuary of Guadalquivir river, Spain (introduced) 37, 6-15 years, 317-701 mm fork length, 690-6,880 g: river Murray, South Australia (introduced) 33, 6 years, 48.8-49.4 cm fork length, 714.5-1,628.4 g: Gölhisar lake, Turkey (introduced) 36, Koi carp: 3-12 years, range 204-700 mm fork length: Lower Waikato region, New Zealand (introduced) 40, 6 years, 360-460 mm fork length, ca 900-1,956 g: Liman lake, Turkey (introduced) 39, 6 years, 475-525 mm fork length, 1,744-2,128 g: Cernek lake, Turkey (introduced) 35, 7 years, 35-36 cm, 629.7-701.4 g: Almus dam lake, Turkey (introduced) 34, 5 years and ca 400-450 mm: Marsh lake, Nebraska, USA (introduced) 41, 5-7 years and ca 650 mm, 12 years and ca 700 mm: Pelican lake, Nebraska, USA (introduced) 41, 483-790 mm, 1,542-7,530 g: Clear lake, Iowa, USA (introduced) 11, 400-700 mm: lake Susan, Minnesota, USA (introduced) 7, females: 6 years, ca 490-660 mm fork length, 7 years, ca 620 mm fork length, males: 6 years, ca 400-600 mm fork length, 7 years, ca 590-680 mm fork length: lake Gariep, South Africa (introduced) 31.
• Observations age and weight FARM: 3 years and 1,340-1,810 g 57, 3+ years, females 1.9-2.8 kg, males 1.1-2.0 kg 55, 2-3 years, females 61.3 cm and 4,350 g, males 63.5 cm and 5,000 g 53, 1,213 g 63.
• Observations TOTAL LENGTH and weight LAB: 490-600 mm and 1.9-6 kg 16, 3 years, 47.2-49 cm, 2,170-2,403.1 g 64, 31.0-33.2 cm, 1,029-1,180 g 24, Koi carp: 22.9-33 cm, 280 g 25, 28.3-37.4 cm, 0.6-1.6 kg 27.

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• Observations WILD: 1:1.2: Liman lake, Turkey (introduced) 39, 1:1: estuary of Guadalquivir river, Spain (introduced) 37, Cernek lake, Turkey (introduced) 35, 1.4:1: Almus dam lake, Turkey (introduced) 34, 1:1.1: lake Gariep, South Africa (introduced) 31, 1.2:1: Amerti reservoir, Ethiopia (introduced) 30.

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• Growth: annulus formation once a year in warm season:
  • Observations annulus formation WILD: April: estuary of Guadalquivir river, Spain (introduced) 37, October-November in scales, November-December in otoliths (warm season): river Murray, South Australia (introduced) 33, Koi carp: annulus formation suspected to take place in late winter (August): Lower Waikato region, New Zealand (introduced) 40.
• Natural growth rate:
  • WILD: back-calculated from scale radius: 104.2 mm fork length in first year, 90.1 mm fork length in second year, 85.1 mm fork length in third year, 72.4 mm fork length in fourth year, 47.4 mm fork length in fifth year, 39.1 mm fork length in sixth year: estuary of Guadalquivir river, Spain (introduced). Comparing with other studies, author found low-growth and high-growth populations depending on latitude: decreasing growth with increasing latitude 37.
  • WILD, Koi carp: ca 100 mm fork length in first three years, ca 70 mm fork length in fourth and fifth year, ca 40 mm fork length in sixth-eighth year, decreasing thereafter: Lower Waikato region, New Zealand (introduced) 40.

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• Observations no difference WILD: estuary of Guadalquivir river, Spain (introduced) 37, Almus dam lake, Turkey (introduced) 34.
• Observations bimodal pattern WILD: males: 6-12 years, 317-678 mm fork length, 690-5,514 g, females: 6-15 years, 383-701 mm fork length, 904-6,880 g: river Murray, South Australia (introduced) 33, females were larger in maximum size than males: lake Gariep, South Africa (introduced) 31.
• Observations beginning of noticeable size difference WILD: Koi carp: males and females similar in length until 6 years of age, from then on, females larger than males: Lower Waikato region, New Zealand (introduced) 40.

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• Growth in polyculture:
  • LAB: JUVENILES (Scaled carp, 126 g) in monoculture or 60-40 or 40-60 polyculture with Blue tilapia (Oreochromis aureus) in 108 L tanks at density 15 IND/tank. After eight weeks, higher specific growth rate (1.02 versus 0.96 under 60-40 versus 0.88 under monoculture), lower FOOD CONVERSION RATIO (1.4 versus 1.6 under 60-40 versus 1.7 under monoculture), and higher mean increase in body weight (76.8% versus 70.5% under 60-40 versus 63.7% under monoculture) in 40-60 polyculture compared to 60-40 polyculture and monoculture (also for Blue tilapia) 65.
  • For polyculture and effects on welfare ➝ D15.
• Growth and noise:
  • LAB: JUVENILES (Scaled carp, 50.5 g) in 147 L glass tanks (100.5 x 40 x 36.5 cm) or a recirculating aquaculture system at density 10 IND/tank, either 80 or 200 lux and either Mozart's "Eine kleine Nachtmusik" or "Romanza - Jeux interdit" (anonymous) during 4 h per day Monday-Friday or no music. Ambient noise level at 123 dB re 1 µPa, music up to 130 dB re 1 µPa. After 106 days, higher weight (161.8 g versus 154.1 g) and higher specific growth rate under Romanza music than control (1.1 versus 1.05%), under 200 lux also higher than under Nachtmusik (and Nachtmusik higher than control) (177 g versus 171.5 g Nachtmusik versus 156.7 g control, 1.18% versus 1.15% Nachtmusik versus 1.07% control). Results indicate beneficial effect of music on growth, but depending on the type of music and more pronounced under 200 than 80 lux 50.
  • LAB: JUVENILES (Koi carp, 3.7 g) in 100 L glass aquaria (70 x 40 x 40 cm) were exposed to urban noise, instrumental music, or Quran performance under water at 66.92±0.16 dB re 1 µPa sound pressure level three times a day for 30 min each. During 35 days, no difference in growth compared to control (57 dB re 1 µPa sound pressure level). After 90 days, higher growth compared to control in the group with the Quran performance (10.7 g versus 8.8 g control) and both instrumental music pieces (9.2-10.1 g). No difference in growth in group with urban noise compared to control (8.6 g versus 8.8 g). Lower FOOD CONVERSION RATIO compared to control in Quran group (1.6 versus 2.2), followed by both instrumental music pieces (1.7-2.1); higher FOOD CONVERSION RATIO compared to control in group with urban noise (2.4 versus 2.2). On day 35, higher specific growth rate compared to control in the Quran group (ca 0.9% versus 0.4%), followed by one of the instrumental pieces (ca 0.5%). Lower specific growth rate compared to control under urban noise (0.1%). On day 90, higher specific growth rate in the Quran group (ca 1.2% versus 1%), followed by one of the experimental pieces (ca 1.1%). No difference between the other instrumental piece and urban noice compared to control (ca 0.9-1%) 26.
• Growth and domestication:
  • LAB: higher relative growth rates (for weight) for heterozygous (scarcely scaled, hump backed; "Mirror carp") than homozygous (fully scaled, torpedo like) JUVENILES (ca 1%/d versus 0.8%/d); no difference in relative growth rate for standard length 20.

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• LAB: four week old FRY in 10 m² tanks at density 250 IND/tank. After eight months, of 671 JUVENILES, 167 (24.9%) with deformities: 37.6% semi-operculum (shortened operculum usually on one side, but also on both sides), 18.2% spinal deformity (mainly lordosis, scoliosis), 17.4% head deformity (larger or smaller head, deformities of jaw), 4.2% stumpbody (shorter trunk, deeper body, semi-operculum, lordosis), 11.4% multiple deformities, 11.2% eye deformities (blindness). Caused slower growth and abnormal swimming (upside down or sideward) 66.

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• LAB: eggs were incubated on dishes in experimental unit with medium containing - among others - MgSO_4 at concentration 0.001, 0.004, 0.007, 0.01, 0.02, or 0.1 mmol/L. At 170 h after fertilisation, 0% mortality and 1-2% deformities under 0.1 and 0.02 mmol/L magnesium. Increasing mortality and deformity rate with decreasing concentration of magnesium up to 100% mortality and 96% deformities under 0.001 mmol/L. Decrease in rate of tail movement before hatching (24 beats/min at 0.1 mmol/L versus 2 beats/min at 0.001 mmol/L) and pigmentation of lateral abdominal skin (relative densities of 26 at 0.1 mmol/L versus 0 at 0.001 mmol/L) with decreasing concentration of magnesium 67.
• LAB: eggs from hormonally stimulated 7-8 kg females were fertilised with milt from hormonally stimulated 5-8 kg males at 22-23 °C and transferred to 20 °C, 24 °C, 28 °C, or 32 °C by changing temperature at 2-3 °C/h. With beginning of exogenous feeding, FRY were transferred to 20 dm³ aquaria at density 400 IND/tank and water changed to 23 °C at rate of 2-3 °C/h. Trace numbers of deformities. Author suspects higher influence of origin of eggs - than temperature - on deformity rate 51.

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• Nest building and substrate: no data found yet.
• Nest building and water velocity: no data found yet.
• Nest building and water depth: no data found yet.
• Nest building: no data found yet.
• For spawning ➝ D21.

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• Courtship sequence:
  • WILD, ADULTS: male smaller in size than female swam slightly below her touching belly with nose: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12.
  • LAB, ADULTS: ripe males and females in earthen ponds. Male followed female for 8-15 s, lagged behind, followed her again, sometimes touching her genital, flank, or cheek with his snout 16.
• Courtship duration: ➝ Courtship sequence.

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• Spawning substrate: submerged vegetation or artifical alternative:
  • Observations WILD, ADULTS: drifting twigs, bottom covered with bog-rush (Juncus), cut-grass (Leersia), smartweed, flag (Scirpus fluviatilis), hornwort (Ceratophyllum), pondweed (Potamogeton): overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, inundated grassland: Danube river 29, among emergent vegetation, mainly giant bulrushes (Scirpus validus) and cattails (Typha spp.): Clear lake, Iowa, USA (introduced) 11.
  • LAB, ADULTS: spawned in earthen pond often close to ribbons of Hyzex sheets (chemically synthesized from polyethylene) to which eggs stuck 16.
• Spawning season: spring-summer to all year:
  • Observations WILD, ADULTS: April-June: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, April-June: estuary of Guadalquivir river, Spain (introduced) 37, females ready to spawn all year long, peak September-April (warm season): various sampling sites, Victoria, Australia (introduced) 32, Koi carp: September-January (spring-summer): Lower Waikato region, New Zealand (introduced) 40, May-June: Clear lake, Iowa, USA (introduced) 11, November (warm season): lake Gariep, South Africa (introduced) 31, throughout the year with peak February-April: Amerti reservoir, Ethiopia (introduced) 30.
• Spawning (day)time:
  • Observations WILD, ADULTS: noon: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12.
  • Observations LAB, ADULTS: spawning period in experimental ponds 06:00-07:17 a.m., 08:30-09:34 a.m., 05:40-06:46 a.m., 07:36-09:50 a.m., 10:48-11:26 a.m. 16.
• Spawning temperature: 15-25 °C:
  • Observations WILD, ADULTS: 15-22 °C: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, 18-19 °C: estuary of Guadalquivir river, Spain (introduced) 37, ≥17 °C: Danube river 29, daily average 16.5-22.6 °C, max 25 °C: various sampling sites, Victoria, Australia (introduced) 32, Koi carp: 16.5 °C: Lower Waikato region, New Zealand (introduced) 40, 18-22 °C: Clear lake, Iowa, USA (introduced) 11, ca 20 °C: lake Gariep, South Africa (introduced) 31, 18.8-23.1 °C: Amerti reservoir, Ethiopia (introduced) 30.
• Spawning salinity: fresh water ➝ D10 D11.
• Spawning and water velocity: no data found yet.
• Spawning depth: 5-165 cm:
  • Observations WILD, ADULTS: 15-90 cm, up to 165 cm: overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, 25-50 cm: Danube river 29, shallow water (mean 1.5-1.6 m): Clear lake, Iowa, USA (introduced) 11.
  • Observations LAB, ADULTS: spawned in 50 cm earthen pond 5-8 cm below surface, but sometimes also 40-46 cm deep 16.
• Spawning density:
  • Observations WILD, ADULTS: several hundred females on <1 acre (ca 4.000 m²): overflowed fields and marshes, lakes along Thompson's lake, Illinois, USA (introduced) 12, large school: Danube river 29.

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• Spawning sequence:
  • LAB, ADULTS: ripe males and females in earthen ponds. Shortly before spawning, quick succession of swimming alongside and nudging the female by the male. Supported each other with pectoral fins, had dorsal fins erect, genitals close to each other. Splashed with caudal fins, released eggs and milt simultaneously. Swam 30-50 cm together, parted 16.
• Spawning duration: no data found yet.

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• LAB, ADULTS: spawners in 1,000 L tanks and temperature gradually increased from 10 to 20 °C at 0.8 °C/d, PHOTOPERIOD from 12 to 14 h either in season (May) or out of season (November). ADULTS were injected with GnRHa and dopamine antagonists to induce spawning. Eggs were collected by stripping, milt via syringe. Shorter latency in spawning after hormone injection in females (13.5 h versus 16.2 h) and higher egg weight (103.8 egg weight/kg body weight versus 58.1 egg weight/kg body weight) in season versus out of season. No difference in spermiation (100%) and spermatozoa motility (86-88%) in males, percentage of ovulation (83%), egg survival to eyed-egg-stage (77.7-81.5%) in females. No difference in percentage of abnormal larvae (1.7-2.1%). Larger size at the beginning of exogenous feeding in larvae reared in season versus out of season (6.8 mm versus 5.3 mm) probably due to later age at beginning of exogenous feeding in season versus out of season (7 versus 5). No difference in lengths 7 days later (6.8-6.9 mm). Results indicate no disadvantage in inducing spawning half a year before spawning season 55.

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• Number of spawns:
  • Observations WILD, ADULTS: 2-3 spawns over 10-14 days: Danube river 29.
  • WILD, ADULTS: based on histological studies, authors concluded that some females spawn once, other multiple times within spawning season: various sampling sites, Victoria, Australia (introduced) 32.
• Fecundity per spawn:
  • Observations absolute fecundity WILD, ADULTS: 1,633-190,778 eggs/female: estuary of Guadalquivir river, Spain (introduced) 37, range annual fecundity 120,000-1,540,000 eggs/female, mean annual fecundity 764,000 eggs/female: various sampling sites, Victoria, Australia (introduced) 32, Koi carp: range 29,800-771,000 eggs/female, mean 299,000 eggs/female: Lower Waikato region, New Zealand (introduced) 40, range 36,955-318,584 eggs/female, mean 170,937 eggs/female: Amerti reservoir, Ethiopia (introduced) 30.
  • Observations absolute fecundity LAB, ADULTS: female of 490 mm, 2.5 kg: 1,500-1,600 eggs (765,000 in total in spawning period) 16.
  • Observations relative fecundity WILD, ADULTS: range annual fecundity 75,000-262,000 eggs/kg, mean annual fecundity 162,000 eggs/kg: various sampling sites, Victoria, Australia (introduced) 32, Koi carp: range 19,300-215,700 eggs/kg body weight, mean 97,200 eggs/kg body weight: Lower Waikato region, New Zealand (introduced) 40, range 98,284-310,864 eggs/kg body weight, mean 177,786 eggs/kg body weight: Amerti reservoir, Ethiopia (introduced) 30.

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• Fecundity and age of spawners:
  • LAB, ADULTS: 2-3 year old spawners were injected with carp pituitary gland hormone. Semen was collected 10 h later via syringe, females were stripped. Higher frequency of motile sperm (75% versus 60%), lower sperm density (17.6 mL x 10^-9 versus 24.6 mL x 10^-9) in 2- compared to 3-year old males. No difference in sperm motility (42-44 s) and spermatocrit (96-97%). Smaller egg diameter (1.1 versus 1.3 mm) and tendency of smaller absolute (523,500 eggs/female versus 657,701 eggs/female) and relative fecundity (123.6 eggs/kg body weight versus 131.4 eggs/kg body weight) in 2- compared to 3-year old females. Combining results for fertilisation and hatching rate, survival rate in LARVAE, length of LARVAE at hatching, and length of LARVAE at beginning of exogenous feeding, tendency of better results in groups with combination of 2 and 3 year old males and 3 year old females 53.

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• For spawning ➝ D1.

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• LAB: JUVENILES (Scaled carp, 116 g) accustomed to pale-blue tanks were transferred to black, white, or green cylindrical 27 L tanks at density 10 IND/tank. After 14 weeks, higher cortisol levels in black- than white-adapted JUVENILES (87.7 ng/mL versus 43.1 ng/mL); JUVENILES in green tanks in between (71.6 ng/mL). Higher total lipids in black- adapted than the other JUVENILES (739.4 mg/100 mL versus 612.8-613.4 mg/100 mL). No difference in glucose, electrolytes, osmolality, haematocrit, triglycerides, cholesterol, liver glycogen 68.
• LAB: JUVENILES in 300 L circular tanks at density 10 IND/tank with either white, black, red, blue, or yellow background colour. After 56 days, highest cortisol levels in red tanks (ca 86 ng/mL). Higher cortisol levels in black than white and yellow tanks (62.1 ng/mL versus ca 40 ng/mL); blue in between (ca 44 ng/mL) 61.

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• LAB: JUVENILES (Scaled carp, 116 g) accustomed to pale-blue tanks were transferred to black, white, or green cylindrical 27 L tanks at density 10 IND/tank. After 14 weeks, no difference in final weights (160.8-165.4 g), but tendency of lower specific growth rate (0.39% versus 0.43%) and higher FOOD CONVERSION RATIO in black- than white-adapted JUVENILES (2.6 versus 2.4); green-adapted in between (0.4%, 2.5) 68.
• LAB: JUVENILES in 300 L circular tanks at density 10 IND/tank with either white, black, red, blue, or yellow background colour. After 56 days, no difference in weight (16.4-18.3 g) and specific growth rate (2.7-2.9%), but lower FOOD CONVERSION RATIO in blue and yellow than red tanks (1.3 versus 1.5); black and white in between (1.3-1.4). Results indicate no effect of contrast between food and background which would be highest in black tanks 61.

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• LAB: JUVENILES in 12 L aquaria were fed red agar-agar pellets laced with amino acids L-cycsteine or L-glutamic acid. Higher consumption of pellets with cysteine (97.3%) compared to glutamic acid (41.8%) and control (17%). Lowest variation in consumption of pellets with cysteine (range 81.8-100%, coefficient of variation 5.1%) compared to glutamic acid (0-88.9%, coefficient of variation 56.4%) and control (0-40%, coefficient of variation 89%). Glutamic acid was preferable to control for 7 of 24 JUVENILES, deterrent for one, and indifferent for 16 JUVENILES. Results indicate individual differences in taste preferences 69.
• LAB: JUVENILES in 12 L aquaria were fed red agar-agar pellets laced with taste substances (5% citric acid = sour, 10% sodium chloride = salty, 10% calcium chloride = bitter, 10% sucrose = sweet) or amino acids. Higher consumption of sour (92.1%) and bitter pellets compared to control (51.7% versus 27.6%). No deterrent pellets. No difference in holding taste pellets in the mouth compared to control. Higher consumption of 6 of 21 amino acid pellets compared to control (33.7-88% versus 14.7%). Highest consumption of pellets with cysteine (99%), alanine (97.2%), proline (51.6%), glutamic acid (36.3%). Pellets with extracts of Chironomids with 100% consumption rate. Lower consumption of 7 amino acids compared to control (0-3.7% versus 14.7%). Lowest consumption, i.e. highest deterrence, of pellets with phenylalanine (1%), serine (1%), valine (0%). Longer time of holding preferred (11.0-18.1 s), shorter time of holding repellent pellets in mouth compared to control (2.9-4.2 s versus 6.6 s in control). In anosmic JUVENILES, no difference in consumption of cysteine pellets (100%) and holding time (14 s) compared to intact individuals. Results indicate that taste is not influenced by olfaction but may be controlled by gustatory receptors in the oral cavity 58.

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• Hearing type:
  • Hearing SPECIALIST 70 71 50.
• Hearing spectrum:
  • LAB: JUVENILES acclimated to 20 °C. Temperature was changed to 15 or 25 °C at rate 1 °C/d. JUVENILES were anaesthetised and secured in plastic tub (45 x 35 x 18 cm) with sandy bottom and were presented with tone bursts at frequencies 0.1, 0.3, 0.5, 0.8, 1, 2, or 4 kHz. Lowest hearing threshold (i.e., best hearing; 57.4-67.3 dB re 1 µPa) and highest sensitivity (4.9-7.8 dB) at 0.3-1 kHz. Lower threshold at 25 °C compared to 15 °C in this range (57.4-62.3 versus 62.8-67.3 dB re 1 µPa). Lower latency of response to single click at 25 °C compared to 15 °C (1.2 versus 1.6 ms) 62.

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• Hearing and categorisation:
  • LAB: ADULTS (Koi carp, 7-11 years, 29-36 cm snout to base of tail) in separated area of home aquarium were presented with music via underwater loudspeaker. Learned to discriminate blues music from classical music from silence via reinforcement (food) to push button when they heard blues. ADULTS also accurately categorised novel music examples into the respective genre. Results indicate higher order auditory processing and capability for categorisation instead of rote memory. In further experiment, ADULTS learned to discriminate music equal in timbre indicating that they used other features for discrimination. In another experiment, ADULTS learned to discriminate classical pieces equal in rhythm but different in melody indicating that they used melody for discrimination 17.

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• LAB: JUVENILES in 3 L bucket were exposed to boat noise (recorded in Danube river and lakes Mondsee and Traunsee, 153 dB), Gaussian noise (by noise generator, 156 dB), and no noise for 30 min. 81% increase in water cortisol under boat noise compared to no noise (ca 0.3 versus 0.2 ng/L water/g body weight), no increase under Gaussian noise. Results indicate higher influence of more fluctuating (in amplitude and frequency) compared to continuous sound 71.
• LAB: JUVENILES (Koi carp, 3.7 g) in 100 L glass aquaria (70 x 40 x 40 cm) were exposed to urban noise, instrumental music, or Quran performance under water at 66.92±0.16 dB re 1 µPa sound pressure level three times a day for 30 min each. During 90 days, erratic swimming when music was first turned on for 2-3 h, more so under sound of urban noise that rapidly increased to maximum sound pressure level compared to the instrumental pieces and the Quran performance with gradual increase 26.

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• LAB: JUVENILES in 50 x 30 x 30 cm tanks with opaque cover on one half and either injected with 5% or 10% acetic acid on upper and lower lips or saline (0.9% NaCl, control). Anomalous behaviour like rubbing the lips on the tank wall, swimming off balance, rocking from side to side in two of the five JUVENILES each in both acid groups at first observation time 30 min after injection (ca 0.3-0.4 events/min). Decreasing over time, completely resolved at last observation time 330 min after injection. No difference in opercular beat rate (gill movement; ca 70-80 beats/min), swimming rate (1-3 swims/min), and use of cover (50-70%) between groups. Results indicate robustness of C. carpio to low concentrations of acid 23.
• LAB: tailfin clips entailed three types of A-fibres that are protected by a myelin sheath for fast conduction of action potentials and transmit well-localised acute pain. Tailfin clips entailed also C-fibres that are protected by glia for slow condution of action potentials and transmit poorly-localised unpleasent dull pain 74.

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• LAB: ADULTS (Koi carp) were injected with either saline (0.9% NaCl), morphine sulfate (5 mg/kg IM), or butorphanol tartrate (10 mg/k IM) and either received anaesthesia (MS-222) or not and either were subjected to unilateral gonadectomy or not.
  Without anaesthesia and surgery, significant decrease in respiratory rate after injection compared to baseline in butorphanol group only. Longer decrease in activity compared to baseline with butorphanol compared to saline (until 18 h post injection versus directly after injection); no change in morphine.
  With anaesthesia but without surgery, larger decrease in respiratory rate until 18 h post recovery from anaesthesia in butorphanol group compared to saline group; no change in morphine group. Longer decrease in activity compared to baseline with saline (12 h versus 6 h under butorphanol versus 3 h under morphine).
  With anaesthesia and surgery, longer decrease in respiratory rate compared to baseline with butorphanol (until last observation time 72 h post recovery versus 24 h with saline versus 18 h with morphine). Decrease in food consumption compared to baseline in butorphanol group directly after recovery, in saline group until 36 h after recovery; no change in morphine group. Compared to injection-only group, longer decrease in food consumption with saline (saline: 36 h post recovery versus 18h with butorphanol and morphine). Longer decrease in activity compared to baseline with morphine (until last observation time 72 h post recovery versus 48 h with saline versus 36 h with butorphanol).
  93% of ADULTS with butorphanol had difficulty with buoyancy and swam with head near the bottom until 48 h after recovery. Results indicate analgesic effect of butorphanol and morphine and advantage of morphine in terms of change in respiratory rate and food consumption rate 25.

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• LAB: JUVENILES (Scaled carp, 116 g) accustomed to pale-blue tanks were transferred to black, white, or green cylindrical 27 L tanks at density 10 IND/tank. After two weeks, JUVENILES in black tanks with dark skin, JUVENILES in white tanks with pale skin, JUVENILES in green tanks with unchanged colour (grey-brownish) 68.

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• WILD: JUVENILES aggregated in May-June, ADULTS aggregated in January-February under ice cover, dispersed from March on, returned to aggregation during pre-spawning in May-June, dispersed thereafter: Clear lake, Iowa, USA (introduced) 11.
• WILD: individuals (420-630 mm, 1,158-3,671 g) stayed in aggregations: warm-water discharge canal, lake Erie, Canada/USA (introduced) 8.
• For aggregation type and daily rhythm ➝ D23.

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• For density in spawning aggregations ➝ D1.

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• LAB: FRY in 30 L aquaria (49 x 29 x 24.6 cm) at densities of 0.2, 0.4, 0.8, 1.6, 3.2, 6.4 IND/L. After ten weeks, decreasing survival with increasing density with 100% at 0.2, 0.4, and 0.8 IND/L and 44.5% at 6.4 IND/L 56.
• LAB: all-male JUVENILES of cross between XX female and homozygous YY male in rectangular tanks of 132 L at density 28.4 kg/m³ (D1) were crowded by shifting movable screens of nylon mesh to achieve densities of 56.8 kg/m³ (D2) or 113.6 kg/m³ (D3). After 15 h, 39 h or 87 h, JUVENILES were immediately sampled or kept in 15 L net for 1 h.
  Cortisol: higher cortisol levels after 15 h crowding in D3 compared to D1 (ca 75 ng/mL versus ca 10 ng/mL), no difference on other observations times or for D2. After additional confinement, increase in cortisol levels in all groups and at all observation times; higher levels compared to D1 at 15 h crowding in D2 (ca 300 ng/mL versus 200 ng/mL) and D3 (ca 250 ng/mL) and at 87 h in D3 (ca 300 ng/mL).
  Free fatty acids: higher plasma free fatty acids compared to D1 after 15 h in D3 (ca 0.5 mM versus 0.3 mM), after 39 h in D2 and D3 (ca 0.4 mM), and after 87 h in D2 (ca 0.4 mM). After additional confinement, increase in free fatty acids compared to no confinement; higher levels compared to D1 at 15 h in D2 (ca 0.6 mM versus 0.3 mM) and D3 (ca 0.7 mM), at 39 h in D2 and D3 (ca 0.4 mM), and after 87 h in D2 and D3 (ca 0.4 mM).
  Glucose: higher glucose levels compared to D1 at 15 h in D2 (ca 3.7 mM versus 2.2 mM), no difference on other observation times or for D3, no effect of confinement.
  Lactate: higher lactate levels compared to D1 at 39 h and 87 h in D3 (ca 3.5-3.7 mM versus 2.7 mM), no effect of confinement.
  Others: lower plasma protein levels compared to D1 at 15 h in D3 (20.8 mg/mL versus 24.1 mg/mL). No effect on haemoglobin and haematocrit 60.
• LAB: all-male JUVENILES of cross between XX female and homozygous YY male in 140 L flow-through tanks at loading densities 0.6 kg/L min (25 IND/tank) or 2.6 kg/L min (100 IND/tank). During 28 days, lower oxygen (53-68% versus 77-91%), lower pH (7.4-7.5 versus 7.6-7.8), and higher conductivity (194-196 µS/cm versus 185-190 µS/cm) in high density compared to low density tanks. Increase in plasma cortisol compared to day 1 on day 3, 8, 14, and 28 (ca 15-60 ng/mL versus 5 ng/mL), higher under high density compared to low density at observation day 3 (ca 60 versus 20 ng/mL). With cortisol in water, higher levels under high compared to low density on almost all observation times (ca 2-5 ng/100 mL versus 0.05-1.5 ng/100 mL), higher compared to day 1 only on day 8 and – under high density – on day 28. Results indicate possibility of measuring cortisol in water instead of plasma which would further eliminate stress (of handling and sampling) for individuals and might be more informative to display chronic stress due to acclimation and thus decrease in plasma cortisol.
  Decrease of free fatty acids in both groups throughout the experiment (0.2-0.3 versus 0.5 mM); no differences between high and low density in glucose (2.3-3.1 mM) and lactate (2.7-4.3 mM) 75.

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• FARM: smaller (65-66 g) or larger (150-152 g) JUVENILES in 500 L tanks of recirculating aquaculture system at either 32 or 64 kg/m³ density. After 30 days, higher specific growth rate (1.5% versus 1.2-1.3%) and lower FOOD CONVERSION RATIO (1.0-1.1 versus 1.3) under 32 kg/m³ than 64 kg/m³ (final densities 51 kg/m³ and 95 kg/m³) 76.
• LAB: FRY in 30 L aquaria (49 x 29 x 24.6 cm) at densities of 0.2, 0.4, 0.8, 1.6, 3.2, 6.4 IND/L. After ten weeks, increasing length and weight with decreasing density with highest length (36.7 mm versus 30.3 mm at 6.4 IND/L) and highest weight (1.9 g versus 0.9 g at 6.4 IND/L) at 0.2 IND/L. Lowest coefficient of variation in length (6.8% versus ca 10-11%) and weight (29.1% versus ca 37-47%) at 0.2 IND/L 56.

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• Managing self-feeder:
  • LAB: JUVENILES learned how to manage a self-feeder (➝ D24) on first day of exposure 15.
• Music categorisation:
  • LAB: ADULTS (Koi carp, 7-11 years, 29-36 cm snout to base of tail) learned via reinforcement (food) to push button when they heard blues music and not push it when there was silence or classical music. Also managed reversal learning where classical instead of blues music was suddenly reinforced. Speed of reversal learning increased with each session 17.
• Presence of food:
  • LAB: JUVENILES learned to associate either of two lights above separate chambers with the presence or absence of food. Time to emerge from shelter (from ca 400 s to 6 s) and search time decreased over run of experiment (from ca 35 s to 15 s). For differences in learning strategies ➝ D5 18.

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• WILD: ADULTS learned to shift home range to mesh bag of cracked corn placed in lake Susan, Minnesota, USA (introduced). Overlap in core area of distribution increased from <10% to ca 70% on the sixth night and almost 100% on the next observation time in the eighth night. No shift during day (➝ D23). Results indicate spatial memory and learning at fast rate 7.

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• Boldness:
  • LAB: JUVENILES from homozygous (fully scaled, torpedo like) and heterozygous (scarcely scaled, hump backed) parents in 130 x 40 x 40 cm tanks. They were categorised as bold or shy based on how fast they left a refuge and how closely they explored a potentially threatening novel object (LEGO Duplo brick). Range in time within one fish length of brick 0-4.2 min. Consistency six weeks later. In a second test, they were assessed on how fast they began feeding under predation risk (open water) and resumed feeding after predation attempt (weight plunging in aquarium). Range in latency to feed 0-10 min, range in latency to resume feeding 0.1-10 min (10 min was cut-off time). Correlation between latency to feed and latency to resume feeding. Negative correlation between "time within one fish length of brick" and both "latency to feed" as well as "latency to resume feeding". Surprising positive correlation between "time within one fish length of brick" and "time within two fish lengths of shoal" (➝ D25) probably due to experimental setup that made compartment with shoal a potentially risky environment and that made swimming to shoal across open water a potentially risky behaviour. No relationship of any behavioural measures with standard length. No relationship between domestication and boldness 20.
  • LAB: JUVENILES were categorised as shy or bold based on how fast they left a shelter after a simulated predator attack (weight plunging in aquarium). Of 745 JUVENILES, 121 entered. No consistency in repetition 14-16 weeks (short term) or 18-23 weeks (long term) later. Lack of entering aquarium possibly due to individual testing which could have stressed the strongly shoaling carp. Negative correlation between "time they spent behind the plants that served as shelters" and "number of grid lines of the aquarium crossed". No correlation with exploration behaviour (➝ D5) 22.
• Risk taking and domestication:
  • LAB: FRY of growth-hormone transgenic Yellow River C. carpio (with higher feeding motivation, more activity and aggression) and size-matched non-transgenic conspecifics in 175 x 35 x 35 cm glass aquaria (water depth 20 cm) divided into five compartments: among them two with feeders separated from buffer zone by transparent glass screen, one with predator Siniperca chuatsi separated by porous transparent glass screen. One feeder dispensed 1 pellet/5 s and the other 2 pellets/5 s. In absence of predator, no difference in feeding rate of transgenic compared to size-matched non-transgenic conspecifics at the two feeders (79% versus 68.7%) and preference of the high-dispensing feeder indicating no difference in feeding competition. In presence of predator, lower feeding rate in general and lower in non-transgenics than transgenics (42.2% versus 59%). At high-dispensing feeder close to compartment with predator, higher feeding rate of transgenics than non-transgenics (5.4% versus 1.1%) indicating higher risk-taking behaviour in transgenic individuals 21.

^{In the structure of menu item 16 and the definition of "SHYNESS-BOLDNESS", we follow 77.}

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• Risk taking and competitive behaviour:
  • LAB: JUVENILES (64.7 mm) were categorised as either risk taking or risk avoiding based on how fast they left a darkened chamber to explore a novel aquarium with food. Range in emergence time 43-843 s with high consistency in the order of emergence over 10 repetitions. No relationship with TOTAL LENGTH. JUVENILES were also categorised as competitive or not based on whether they were first at a feeding tube. Displayed no direct aggression. Consistency in the order of feeding at the tube over 10 repetitions. No relationship with TOTAL LENGTH. Positive relationship between risk taking and competitive behaviour.
    Experiment 2: JUVENILES (1 year, 25.3 g) were categorised as either risk taking or risk avoiding. Individual differences, but in general, higher resting metabolic rate in risk takers compared to risk avoiders (ca 117 versus 101 mg O2/kg/h) indicating higher maintenance costs and fitting with model of higher activity in risk takers.
    Experiment 3: JUVENILES (1 year, 64.7 g) categorised as risk taking or risk avoiding in 60 L tanks at density 20 IND/tank. After 10 weeks, higher levels of glucose (ca 150 mg/dL versus 90 mg/dL) and lactate (ca 55 mg/dL versus 28 mg/dL) in risk avoiders than risk takers.
    Necessity to test JUVENILES in groups because they showed signs of distress when isolated may have influenced results in that individuals feel more secure in groups or are affected by their conspecifics' responses 19.
• Exploration:
  • LAB: JUVENILES from homozygous (fully scaled, torpedo like) and heterozygous (scarcely scaled, hump backed) parents in 130 x 40 x 40 cm tanks. Were categorised as either explorative or avoiding based on how fast they left a refuge to explore a novel aquarium. Range in emergence time 0-60 min (60 min was cut-off time), range in time spent in open arena 0-9.3 min, range in time spent in upper water zone 0-0.9 min. Consistency in rank order in latency to emerge in second trial six weeks later. Correlation between the three measures. No relationship of any behavioural measures with standard length 20.
  • LAB: JUVENILES were categorised as explorative or avoiding based on how fast they left a darkened chamber to explore a novel environment. Of 745 JUVENILES, 121 entered. No consistency in repetition 14-16 weeks (short term) or 18-23 weeks (long term) later (with altered gravel, plants, wall colours to ensure novelty of habitat). Lack of entering aquarium possibly due to individual testing which could have stressed the strongly shoaling carp 22.
• Proactivity and reinforcement learning:
  • LAB: JUVENILES were categorised as either proactive or reactive based on how fast they left a darkened chamber to explore a novel aquarium with food. High consistency over three repetitions. No relationship with TOTAL LENGTH. JUVENILES learned to associate either of two lights above separate chambers with the presence or absence of food, but in two different ways. Majority of reactive JUVENILES learned to discriminate light colours, majority of proactive JUVENILES chose one of chambers randomly and switched to other one if no food present. Proactive JUVENILES that learned to discriminate light colours took more trials than reactive JUVENILES (ca 21 versus 15) 18.

^{In the structure of menu item 16 and the definition of "EXPLORATION-AVOIDANCE", we follow 77.}

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• LAB: JUVENILES were categorised as passive or active based on how many grid lines of the aquarium they crossed (passive: 0-26, active: 100-338) and how long they spent behind the plants that served as shelter (passive: 205-600 s, active: 0-327 s) in a study assessing reactions to a simulated predator attack (➝ D4). Size-matched pairs of similar behavioural types (both active or both passive) were placed in a different aquarium and observed. Passive JUVENILES spent more time in shelter than active ones (ca 400 s versus 300 s), but no difference in activity. Negative correlation between activity and time spent in shelter. Consistency in repetition two days later. After five months, active JUVENILES with deeper body than passive ones 22.

^{In the structure of menu item 16 and the definition of "ACTIVITY", we follow 77.}

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• LAB: JUVENILES from homozygous (fully scaled, torpedo like) and heterozygous (scarcely scaled, hump backed) parents in 130 x 40 x 40 cm tanks. They were categorised as sociable based on their shoaling tendency, i.e. how close they swam to conspecifics. Range in time within two fish lengths of shoal 0-8.8 min. Consistency six weeks later. Surprising positive correlation between "time within one fish length of brick" (➝ D4) and "time within two fish lengths of shoal" probably due to experimental setup that made compartment with shoal a potentially risky environment and that made swimming to shoal across open water a potentially risky behaviour. No relationship of any behavioural measures with standard length 20.
• For isolation stress ➝ D5.

^{In the structure of menu item 16 and the definition of "SOCIABILITY", we follow 77.}

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• LAB: JUVENILES were not tested in isolation due to strong signs of stress and instead tested in groups (➝ D5) 19.
• LAB: JUVENILES were tested in pairs to avoid isolation stress (➝ D5) 18.

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• Air exposure:
  • LAB: JUVENILES (479 g) were transferred from pond to tank and kept either 2 min or 5 min in the air in a net. Lower cortisol levels (ca 550 µmol/L versus 950 µmol/L) and higher glucose levels in 2 min- and 5 min-groups compared to individuals directly transferred to tank (ca 7.2 mmol/L versus 5 mmol/L). Negative correlation between cortisol and glucose.
    JUVENILES (672 g) were transferred from pond to tank that either contained pond water or pond water and anaesthetic Menocain at 0.2 g/L. No difference in cortisol levels, but lower glucose levels in anaesthetised individuals (ca 4.7 mmol/L versus 6.5 mmol/L). Results indicate potential of anaesthetic to decrease stress response by preventing depletion of energy resources 57.

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• LAB: all-male JUVENILES of cross between a female and a homozygous YY male were placed in 5 L nets at density 5 IND/net in confinement tank for 0.5, 1, or 3 h and then returned to holding tank. Similar increase in cortisol at all time points compared to control (ca 250-330 ng/mL versus ca 40 ng/mL). Back at pre-stress levels at first observation time 1 h after confinement. Glucose levels increased with increasing confinement period, higher at all times than control (ca 4.5-5.5 mM versus 2 mM). Glucose increased still after return to holding tank until at least 4 h post confinement. Back at pre-stress levels at next observation time 22 h after stress. Decrease in plasma lactate levels compared to control at 3 h confinement and at 1 h post stress, thereafter increased to pre-stress levels. Increase in plasma free fatty acid levels compared to control at 3 h confinement. Remained increased until at least 22 h post stress. Decrease in haemoglobin levels compared to control at 0.5 h and 3 h of confinement as well as 1 h post stress (10.6-10.7 mg/dL versus 12.3 mg/dL). No effect on cholesterol and triglycerides 59.
• LAB: all-male JUVENILES of cross between XX female and homozygous YY male in 140 L tanks accustomed to 20 g feed/kg^0.8/d (ca 3-4% body weight) were a) switched to low feeding regime (5 g feed/kg^0.8/d, ca 0.7-1% body weight) for 80 days, brought back to accustomed feeding for 22 days (L->H) or b) kept at feeding for 22 days, then switched to 5 g feed/kg^0.8/d for 80 days (H->L). JUVENILES were immediately sampled or placed in 5 L nets at density 5 IND/net in confinement tank for 0.5, 1, or 3 h and then returned to holding tank. Before confinement, higher triglycerides in L->H than H->L group (2.3 mM versus 1.1 mM); no difference in cortisol (30.5-33 nM), glucose (1.5-3 mM), lactate (2.7-3.2 mM), free fatty acids (0.1-0.2 mM), cholesterol (3.5-4 mM). Confinement increased cortisol compared to control, more so in L->H individuals at 0.5 h confinement (ca 650 versus 450 nM); no difference at 1 h and 3 h. Back to pre-stress levels at next observation time 3 h post stress. Confinement increased glucose compared to control faster in L->H group (ca 5-6 mM versus 3 mM) and higher. Higher glucose level than H->L group at 0.5 and 1 h (ca 5 mM versus 2.5-3 mM) and at 3 h after confinement (ca 6 mM versus 4 mM). Higher glucose in H->L group compared to control at 3 h confinement and 3 h post stress (ca 4 mM versus 1.5 mM). Back to pre-stress levels at next oberservation time 21 h post stress. Higher free fatty acids compared to control at 3 h confinement and at least until 21 h post stress (ca 0.4-0.6 mM versus 0.1-0.2 mM). Results indicate advantage of fasting before stressful events 78.
• For confinement after long-term crowding ➝ D26.

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• FARM: 3 year old ADULTS were transported in long-distance truck with water oxygenated from oxygen container in two batches at 300 kg/m³ for 10 h. In first transport, decrease in cortisol levels (1,242 µmol/L versus 803 µmol/L) and increase in glucose levels after transport compared to at the beginning (7.2 mmol/L versus 4.4 mmol/L). In second transport, where individuals had been caught feeding from production pond at 20 °C directly into truck with 16% oxygen saturation an 15 °C, majority was found dead after 10 h. No difference in cortisol levels (245.4-346 µmol/L) and increase in glucose levels compared to beginning of transport (31.8 mmol/L versus 6.8 mmol/l with chyme remnants versus 9.9 mmol/L with heavily filled digestive tract).
  2 year old JUVENILES were transported in truck with water aerated by truck compressor at 250 kg/m³ for 2 h. No difference in cortisol levels (308.2-382.7 µmol/L), but increase in glucose levels compared to beginning of transport (7.6 mmol/L versus 5.4 mmol/L). Results indicate importance of empty stomach, avoidance of stress before transport, and similarity in water conditions between transport tank and previous water body 57.
• LAB: ADULTS were transported in 3.3 m³ tank at density 364 kg/m³ for 12.5 h. No increase of cortisol levels compared to loading before transport (258.6-301.9 ng/mL). Decrease of glucose after 7 h of transport (5.9 mmol/L versus 10.8 mmol/L), but no difference after 12.5 h (10.1 mmol/L). Increase of ammonia (ca 450 µmol/L versus 260 µmol/L), lactate (9.2 mmol/L versus 4.8-7.9 mmol/L), lactate dehydrogenase (ca 13.5 µkat/L versus 9.5 µkat/L), aspartate aminotransferase (ca 3.5 µkat/L versus 1.7 µkat/L), creatine kinase (ca 770 µkat/L versus 470 µkat/L), metamyelocyte count (ca 6.5 g/L versus 3.5 g/L) and decrease of alanine aminotransferase (ALT; 0.1 µkat/L versus 1.2-1.6 µkat/L) as well as total proteins (29 g/L versus 33 g/L) after 12.5 h of transport 64.
• LAB: individuals were netted in farm and transported for 20 min in 1 m³ tank to laboratory. Higher cortisol levels after transport than after netting (ca 151 ng/mL versus 75 ng/mL) 79.

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• Polyculture:
  • LAB: JUVENILES (Scaled carp, 126 g) in monoculture or 60-40 or 40-60 polyculture with Blue tilapia (Oreochromis aureus) in 108 L tanks at density 15 IND/tank. After eight weeks, higher haematocrit (43.8 versus 37.2-40.2 ng/mL), cholesterol (134.5 versus 111.9-121.3 mg/100 mL), and triglycerides (170.7 versus 144.1-148.2 mg/100 mL) under mono- than polyculture. No difference in cortisol (56.6-83 ng/mL), glucose (82.7-95.5 mg/100 mL), osmolality (0.27 Osmol/kg). Schooling behaviour under monoculture was disrupted in polyculture 65.

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• Stunning rules: to minimise pain reactions and enhance welfare before slaughter:
  1. induce insensibility as fast as possible,
  2. prevent recovery from stunning,
  3. monitor effectiveness (observations, neurophysiological measurements) 80.

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• Comparison of stunning methods:
  • FARM: among 17 stunning processes in farms in Germany, 10 processes entailed electrical plus percussive stunning, four entailed electrical stunning, three percussive stunning. For details of electrical stunning protocols ➝ 27. Higher percentage of loss of behavioural indicators of consciousness (92.6% versus 71.9% versus 69.2% of individuals) and lower percentage of individuals showing signs of consciousness at time of slaughter after electrical plus percussive stunning than after electrical or percussive stunning (11.8% versus 28.1% versus 46.2%). Higher percentage of problems with percussive than electrical stunning (23.1% injuries from mishits versus 12.5% injuries from contact with electrodes). Among 17 processes, killing (at 0.5-3 min after stunning) in 10 cases by evisceration, in 7 cases by gill cut or destruction of the heart 27.
  • LAB: ADULTS were stunned either by immersion in clove oil (1 mL/L) or CO2, hypothermia (placement in ice water of 0.6-1.5 °C), or asphyxia.
    Under CO2, aversive behaviour, tried not to open mouths and operculum. After 185 s, loss of equlibrium, violent behaviour. Aversive behaviour until 447 s. After 500 s, no reflexes. After 915 s, loss of opercular movement. Other observations: increase in mucus production, darkening of gills.
    Under hypothermia, slow swimming until 675 s. Occasional erratic swimming, tremors. After 967 s, loss of equilibrium and movements. After 1,974 s, no reflexes. After 2,932 s, loss of opercular movement. Other observations: gasping for air.
    Under asphyxia, violent behaviour. After 293 min, loss of opercular movement.
    Under clove oil, calm swimming, loss of equilibrium after 65 s. After 125 s, loss of reflexes. After 225 s, loss of opercular movement. Later peak of rigor mortis under clove oil compared to other methods (at observation time 60 h after death versus 36 h under hypothermia versus 12 h under CO2 versus 3 h under asphyxia) probably due to fewer muscle contractions. Results indicate most rapid and least irritating stunning by immersion in clove oil 24.
• Electrical stunning:
  • LAB: ADULTS were restrained in U-shaped adjustable PVC grid or fenestrated aquarium and electrically stunned with either 50 V (sinusoidal a.c.) and 50 Hz for 5 min (ca 0.1 A/dm² current density) or 150 V and 50 Hz for 1 min (0.3-0.4 A/dm² current density). No recovery in 4 of 32 individuals. With 5 min stun, return of opercular movement after 2-5 min, with 1 min stun, after 0.8-7 min. Afterwards, return of vestibulo-ocular reflex after 1-25 min, attempt of righting behaviour in 19 individuals after 4-55 min. 30 s after stunning, visually evoked responses in 31 out of 32 individuals indicating consciousness. The results hint that in carp, absence of behavioural indicators is not necessarily connected with unconsciousness 27.

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• FARM: among 17 stunning processes in farms in Germany, large variation in plasma cortisol (62.1-337.7 ng/mL), glucose (2.7-16.6 mmol/L), haematocrit (25.5-41.8%), and lactate levels (1.0-41.8 mmol/L) probably due to pre-slaughter handling. Higher cortisol levels (228.3 ng/mL versus 114.0 ng/mL versus 151.3 ng/mL) and lower haematocrit levels with percussive compared to electrical stunning or combination of electrical and percussive stunning (27.4% versus 34.4% versus 34.7%). Higher glucose levels with percussive or a combination of electrical and percussive stunning compared to electrical stunning (6.7 mmol/L versus 4.2 mmol/L versus 3.2 mmol/L) 27.
• LAB: individuals were netted in farm, transported for 20 min in 1 m³ tank to laboratory, and stunned by either percussion, chilling in ice slurry, or asphyxiation in CO2. Higher cortisol levels with live chilling than with percussive stunning (ca 280 ng/mL versus 100 ng/mL), CO2 asphyxiation in between (ca 210 ng/mL) 79.
• LAB: ADULTS were stunned either by percussion (blow to the head with a hammer), electricity, or asphyxia for 90 min. For electrical stunning, two electrodes were placed on the skull and operated with 4.5 mA (DC) and ca 300 V for 3 s once or twice until beginning of seizure. Higher cortisol and higher glucose levels in asphyxia (cortisol range: 633.7-800+ ng/mL, glucose average: 11.4 mmol/L) compared to electrical stunning (cortisol range: 3.2-800+ ng/mL, glucose average: 6.1 mmol/L) compared to percussive stunning (cortisol range: 0.5-676.5 ng/mL, glucose average: 3.9 mmol/L) 63.
• For stunning methods and aversive behaviour ➝ D34.

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